Family Tetillidae Sollas, 1886
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Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest © Kluwer Academic/Plenum Publishers, New York, 2002 Family Tetillidae Sollas, 1886 Rob W.M. Van Soest1 & Klaus Rutzler2 'Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands, ([email protected]) ^ National Museum of Natural History, Smithsonian Institution, Washington DC. 20560, U.S.A. ([email protected]) Tetillidae Sollas (Demospongiae, Spirophorida) are for the most part globular sponges ('golf ball' sponges) belonging to a small order on account of their possession of sigmaspire microscleres and triaene megascleres. The characteristic megasclere is the protriaene, which is found in most members of the family with very few exceptions. In addition there are long oxeas and frequently anatriaenes, rarely also calthrops or amphitriaenes. The skeletal architecture is strictly radial. Eight valid genera are recognized, differentiated by presence of cor- tical structures, specialized pore-sieves, and composition of the spicule complement. Tetillids are common and ubiquitous sponges in all oceans and at all depths. Keywords: Porifera; Demospongiae; Spirophorida; Tetillidae; Acanthotetilla; Amphitethya; Cinachyra; Cinachyrella; Craniella; Fangophilina; Paratetilla; Tetilla. DEFINITION, DIAGNOSIS, SCOPE megascleres). The family comprises more than one hundred species distributed over all oceans and habitats. Synonymy Biology [Lophurellidae] Gray, 1872a. [Casuladae] Gray, 1872a. [Tethyina] Carter, 1875b (not Tethyadae Gray, 1848). Tetillidae Tetillids have a preference for sedimented habitats and some Sollas, 1886a. Tethyopsillidae Lendenfeld, 1903. Tethydae species possess a root of long spicule bundles to attach them to the Lendenfeld, 1907. Ectyonillidae Ferrer-Hernandez, 1914a. substrate. Reproductive patterns range from the extrusion of fertilized Craniellidae de Laubenfels, 1936a. eggs, which fix to the substrate and develop directly, to incubation of complete young sponges which are then expelled by localised Definition breakdown of the pinacoderm. No free larvae have yet been described. Spirophorida with sigmaspire microscleres and a radiate skeleton of triaenes and oxeas. Remarks Diagnosis Gray (1867a) included sponges of this group under the genus name Tethya in a larger Tethyadae comprising a wide range of Typically with spherical growth form, often with characteristic tetractinomorph genera. Carter (1875b) was the first to distinguish pits called porocalices containing the inhalant and occasionally also a separate group (subfamily) for tetillids, but like Gray called the exhalant orifices. Cortical region often strengthened by collagen group Tethyina. The explanation for this is that at that time there fibers and special cortical megascleres, but this region may be thin was still a great confusion over the identity of Tethya. Lamarck or absent in several genera. Skeleton with tetraxonic and monaxonic (1815) included in his genus Tethya next to T lyncurium megascleres (triaenes, huge oxeas) organized in a radiate pattern of ( = T aurantium) and others, Tethya cranium (Miiller, 1789 as spicule bundles, often spiralling outwards from the centre of the Alcyonium). Carter was aware of the great difference between the body; oxeas, protriaenes and anatriaenes are most common and type of Tethya (i.e., T lyncurium) and T cranium (see for example these often protrude from the surface producing a conulose or hairy Carter, 1872a), but he ignored Schmidt's (1870) proposal to surface. In sponges provided with porocalices they form a palisade employ a separate genus Craniella for Alcyonium cranium. of long spicules, protruding far beyond the sponge surface, sur- Schmidt (1870) included Craniella in his family Ancorinidae along rounding the porefields or oscular apertures, which lie in rounded with Stelletta and other astrophorid sponges, but Sollas (1886a) depressions. Microscleres are unique contorted microspined sig- revived Carter's Tethyina under a different name by erecting the maspires. Reproduction is oviparous without a larval stage, or family Tetillidae based on Schmidt's (1868) genus Tetilla. In many viviparous with production of young adults within a parent. ways this was an unwise choice of type genus, inspired by some notion that Tetilla would represent the most primitive of tetillid Scope genera. The description of Tetilla euplocamos by Schmidt (1868: 40, pi. V fig. 10) is insufficient to conclude with certainty what its Out of twenty six nominal genera eight are considered valid, properties were. Selenka (1880: 469) thought to recognize Acanthotetilla, Amphitethya, Cinachyra, Cinachyrella, Craniella, Schmidt's species in an intertidal habitat in the Bay of Rio de Fangophilina, Paratetilla, Tetilla. They are differentiated on gross Janeiro, and provided some additional data. Sollas (1888) was able morphological structure (presence or absence of a cortical region, to verify from a Schmidt slide that at least protriaenes were pres- presence or absence of specialized pore areas), and spicule comple- ent, but apparently this species has no sigmaspires (cf. below). ment (possession of calthrops, amphitriaenes or other auxiliary Even though protriaenes are of common occurrence in tetillids, 85 86 Porifera • Demospongiae • Spirophorida • Tetillidae they are by no means confined to that group and we have to rely and Tethyopsillidae was incorrect, since the type of Tetillidae and on the subtle tetilliform protriaene-type as proof that Tetilla genus Tetilla (T euplocamos) has no sigmaspires and thus would euplocamos is a member of what is generally understood as Tetilla belong in Tethyopsillidae, making this family an objective synonym and the Tetillidae. In summary: we have a type species of the type of Tetillidae. De Laubenfels (1936a) for the same reason refused to genus of a family lacking the major synapomorphy of the order to accept Tetilla euplocamos as a close relative of Craniella, and which the family belongs! Some confidence may be obtained from erected a separate family Craniellidae for the genera possessing sig- the fact that Tetilla species lacking sigmaspires are not uncommon. maspires. Tetilla and Tetillidae were defined as 'Choristida lacking Despite this dubious type species designation, Sollas' (1888) mono- microscleres' and an unrelated group of genera was assigned to it. graphic treatment of Tetillidae was followed by most subsequent Needless to say that neither scheme gained general support. authors with the exception of Lendenfeld (1888, 1903, 1907) and de Laubenfels (1936a). Lendenfeld continued to employ Tethya for Previous reviews sponges answering to the definition of Craniella, but at least kept this in the family Tetillidae. He erected a family Tethyopsillidae for Sollas (1888), Lendenfeld (1903), Wilson (1925), Topsent sponges without sigmaspires. Lendenfeld's division into Tetillidae (1928c), Levi (1973), Rutzler (1987). KEY TO GENERA (1) Specialized pore-bearing pits ('porocalices') present 2 No porocalices 6 (2) Spicules include medium-sized heavily spined oxeas ('megacanthoxeas') Acanthotetilla No megacanthoxeas 3 (3) Among the megascleres there are short-shafted triaenes with equal-sized rays (calthrops). N.B. these may have bifid cladi but are not to be confused with long-shafted amphiclads Paratetilla No calthrops, all triaenes - if present - are long-shafted 4 (4) In cross section, there is a clear cortical region with special cortical megascleres (short oxeas) and collagenous reinforcement Cinachyra In cross section there is no discernible peripheral layer different from that of the interior 5 (5) Porocalices all similar, undifferentiated morphologically Cinachyrella Porocalices differentiated into a morphologically distinct inhalant and exhalant type Fangophilina (6) Stalked sponges with differently structured stalk and main body; among the megascleres of the stalk there are triaenes with cladi on both ends (amphitriaenes, -diaenes, -monaenes, collectively called amphiclads) Amphitethya No amphiclads, if stalked, then it is a thin root-like projection, not a proper stiff stalk 7 (7) In cross section, there is a clear cortical region with collageneous reinforcement Craniella In cross section there is no discernible peripheral layer different from that of the interior Tetilla ACANTHOTETILLA BURTON, 1959 Previous review Synonymy Van Soest (1977b). Acanthotetilla Burton, 1959a: 201. Acanthocinachyra Levi, Description of type species 1964b: 386. Acanthotetilla hemisphaerica Burton, 1959a (Fig. 1A-B) Type species Synonymy. Acanthotetilla hemisphaerica Burton, 1959a: 201, fig. 5; Van Soest, 1977b: 2, pi. I figs a-b, pi. II figs c-d, text-fig. 1. Acanthotetilla hemisphaerica Burton, 1959a: 201 (by Material examined. Holotype: BMNH 1936:3:4:530 - John monotypy). Murray Exped. stat. 45, coast of S Arabia, litothamnion bottom, 38 m depth. Definition Description. Semiglobular (Fig. 1 A), 4—5 cm diameter. Colour in alcohol yellowish brown. Surface hispid-bristly due to megascle- Tetillidae with megacanthoxeas as auxiliary megascleres. res projecting 1 mm or more beyond the ectosome. Porocalices numerous, 1-2 mm diameter, scattered over the surface. No appar- Diagnosis ent oscules. Consistency hard, incompressible. Skeleton radiate, forming a tight palisade at the surface. The main megascleres are Globular, hemispherical or irregularly massive sponges. smooth long oxeas and medium-sized megacanthoxeas, but protri- Surface extremely hispid and